Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 125(3): 587-608. November 2019

RECENT BRACHIOPODS FROM THE TONGA ISLANDS, SW PACIFIC: TAXONOMY AND BIOGEOGRAPHY

MARIA ALEKSANDRA BITNER

Institute of Paleobiology, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warszawa, Poland. E-mail: [email protected]

To cite this article: Bitner M.A. (2019) - Recent brachiopods from the Tonga Islands, SW Pacific: taxonomy and biogeography. Riv. It. Paleont. Strat., 125(3): 587-608.

Keywords: Brachiopoda; systematics; biodiversity; Pacific archipelagos;B ordau 2.

Abstract. Twenty species of Recent brachiopods belonging to the genera Neoancistrocrania, Basiliola, Basiliolella, Dyscolia, Abyssothyris, Xenobrochus, Terebratulina, Fallax, Septicollarina, Frenulina, Amphithyris, Annuloplatidia, Leptothyrella, Dallina, Campages, Thecidellina and Minutella have been identified in the material collected during the French cruise Bordau 2 to the Tonga Islands, South-West Pacific. Apart from Frenulina sanguinolenta all species represent the first records for the Tonga Islands. The investigated brachiopod fauna shows the greatest affinity to that from Fiji and New Caledonia, having 16 and 12 species in common, respectively. Although less affinity is observed with the New Zealand fauna, there are two species, Terebratulina australis and Amphithyris buckmani reported so far only from New Zealand, Fiji and Tonga. The biodiversity of brachiopods in Tonga is similar to that in Fiji but half as great as that in New Caledonia and New Zealand regions and much higher than in French Polynesia.

Introduction origin, the eastern non-volcanic coral limestone. So far only two species, Novocrania turbinata (Poli, 1795) Since 1976 the South-West Pacific region has and Frenulina sanguinolenta (Gmelin, 1791) have been been intensively surveyed within the program Trop- reported from Tonga (Saito & Endo 2001; Logan ical Deep-Sea Benthos (formerly MUSORSTOM) 2007; Robinson 2017). Although both species are established by the Muséum national d’Histoire na- well known and widely distributed in the western turelle (Paris, France) and the Institut de Recher- Pacific, in the studied material they are either absent che pour le Développement (Nouméa, New Cale- (N. turbinata), or very rare (F. sanguinolenta). How- donia). The present report focuses on the Tonga ever, Robinson (2017) recognized N. turbinata in the island group, explored during the cruise Bordau 2 University of Tokyo collection from Tonga. Among in 2000. The name of the cruise refers to the BOR- the 20 species identified in the studied material, 19 Der of the Indo-AUstralian plate (Bouchet et al. are reported for the first time from the Tonga Is- 2008). Tonga, comprising more than 170 islands, lands. is an archipelago in the South Pacific Ocean close to Fiji, about two-thirds of the way from Hawaii to New Zealand (Fig. 1). It lies along the boundary of the Pacific and Indian-Australian tectonic plates Material and methods and consists of two geologically different, parallel The material presented here was collected during the cruise chains of islands; the western islands are of volcanic Bordau 2 (https://expeditions.mnhn.fr/campaign/bordau2) to the Tonga Islands, SW Pacific (Fig. 1). The expedition was organized by the Muséum national d’Histoire naturelle, Paris and by the Institut Received: March 12, 2019; accepted: May 17, 2019 de la Recherche pour le Développement, Nouméa, New Caledonia 588 Bitner M.A.

Class Craniata Williams, Carlson, Brunton, Holmer & Popov, 1996 Order Craniida Waagen, 1885 Superfamily Cranioidea Menke, 1828 Family Craniidae Menke, 1828 Genus Neoancistrocrania Laurin, 1992 Type species - Neoancistrocrania norfolki Laurin, 1992, by original designation of Laurin (1992: 344)

Neoancistrocrania norfolki Laurin, 1992 Fig. 2A-C

1992 Neoancistrocrania norfolki Laurin, pp. 344-346, pl. 1, figs. 1-6, pl. 2, figs. 1-6. 1997 Neoancistrocrania norfolki - Laurin, pp. 417-418, fig. 41A-E. 2009 Neoancistrocrania norfolki - Bitner, p. 6, fig. 2A-F. 2010 Neoancistrocrania norfolki - Zezina, p. 1179. 2014a Neoancistrocrania norfolki - Robinson, fig. 3A-H. 2014b Neoancistrocrania norfolki - Robinson, p. 542, fig. 2A-E. 2015 Neoancistrocrania norfolki - Bitner, p. 35, fig. 2A-B.

Material examined: Tonga, Bordau 2 cruise, stn DW 1535, one dorsal valve (MNHN IB-2013-678); stn DW 1602, one dorsal valve (MNHN IB-2013-679); stn DW 1605, one ventral valve, broken (MNHN IB-2013-680). Depth range: 263-441 m. Measurements: Max. length 14.8 mm, width 20.8 mm.

Remarks. Neoancistrocrania norfolki is very rare in the studied material, found in only 3 stations. Morphologically this species is readily distinguished Fig. 1 - Map of Tonga Islands, showing location of the brachio- from Novocrania Lee & Brunton, 2001 by its massive, pod-bearing stations. mineralized ventral valve and two erect divergent processes on the dorsal valve interior (Laurin 1992, on R.V. “Alis” and carried out from 31 May 2000 to 22 June 2000. 1997; Bitner 2009, 2015) and by unique features of Brachiopods were found in 69 out of 138 stations. The exact loca- tion, depth and species identified at each stations are given in the the soft tissues (Robinson 2014a, 2014b). However, Appendix. Although the collection is rich overall (1212 specimens), molecular analysis shows that “Neoancistrocrania is nine species are rare (3 or fewer specimens). no more distantly related to the Northern clade of Samples were collected using a Warén dredge (DW) or a Novocrania than are other Novocrania clades” (Cohen trawl (CP, CH). To remove soft tissues, specimens were treated with hypochlorite bleach, followed by a water wash. For scanning elec- et al. 2014: 145) suggesting generic synonymy of tron microscope (SEM) examination, the selected specimens were Neoancistrocrania under Novocrania. Such conflict be- mounted on stubs, coated with platinum, and investigated using a tween the morpho-classification and molecular sys- Philips XL-20 microscope at the SEM laboratory of the Institute of tematics is observed in other brachiopod groups as Paleobiology, Warszawa. All the material is deposited in the collection of the Museum national d’Histoire naturelle, Paris, France under the well (Cohen & Bitner 2013; Bitner & Cohen 2015). catalogue number MNHN IB-2009-516 to IB-2009-562, IB-2009- Previously N. norfolki had been found only in 1198, IB-2009-1240, IB-2013-678 to IB-2013-790. the West Pacific (Laurin 1997; Bitner 2009, 2015; Cohen et al. 2008, 2014). The present finding ex- tends its geographical range eastward. Systematic part

Subphylum RHYNCHONELLIFORMEA Phylum BRACHIOPODA Duméril, 1805 Williams, Carlson, Brunton, Holmer & Popov, 1996 Subphylum CRANIIFORMEA Popov, Bassett, Class Rhynchonellata Williams, Carlson, Holmer & Laurie, 1993 Brunton, Holmer & Popov, 1996 Recent brachiopods from the Tonga Islands, SW Pacific 589

Fig. 2 - A-C - Neoancistrocrania norfolki Laurin, 1992, Tonga, Bordau 2, stn DW 1602, 263-320 m, outer, inner and tilted views of dorsal valve, MNHN IB-2013-679. D-H - Basiliola lucida (Gould, 1862); (D-E) inner and oblique views of posterior part of dorsal valve, MNHN IB-2013-700, SEM, stn DW 1635, 320-323 m; (F-H) dorsal, lateral and anterior views of articulated specimen, MNHN IB-2013-699, stn DW 1587, 309-400 m. I-M - Basiliola beecheri (Dall, 1895), Bordau 2; (I-K) dorsal, lateral and anterior views of articulated specimen, MHNH IB-2013-694, stn DW 1619, 591-593 m; (L-M) inner and oblique views of posterior part of dorsal valve, MNHN IB-2013- 683, SEM, stn CP 1545, 444-447 m. N-S - Basiliolella colurnus (Hedley, 1905), Bordau 2; (N-Q) dorsal, lateral views and dorsal, anterior views of two articulated specimens, MNHN IB-2013-706, stn CP 1545, 444-447 m; (R-S) inner and oblique views of posterior part of dorsal valve, MNHN IB-2013-703, SEM, stn DW 1536, 320-323 m. 590 Bitner M.A.

Order Rhynchonellida Kuhn, 1949 1895 Hemithyris beecheri Dall, p. 717, pl. 31, figs. 1-4. 2008 Basiliola beecheri - Bitner, pp. 427-428, fig. 5H-L cum( syn.). Superfamily Pugnacoidea Rzhonsnitskaya, 1956 2009 Basiliola beecheri - Bitner, p. 6-7, fig. 3D, E. Family Basiliolidae Cooper, 1959 2010 Basiliola beecheri - Zezina, p. 1179. Genus Basiliola Dall, 1908 2015 Basiliola beecheri - Bitner, pp. 35-36, fig. 2G-H. Type species - Hemithyris beecheri Dall, 1895, by original designation of Material examined: Tonga, Bordau 2 cruise, stn DW Dall (1908: 442) 1509, one articulated specimen (MNHN IB-2009-544); stn CP 1510, 7 articulated specimens (MNHN IB-2013-681); stn DW Basiliola lucida (Gould, 1862) 1538, 2 articulated specimens (MNHN IB-2009-545); stn CP 1539, 2 articulated specimens (MNHN IB-2009-542); stn DW 1543, one Fig. 2D-H articulated specimen (MNHN IB-2013-682); stn DW 1544, one articulated specimen (MNHN IB-2009-540); stn CP 1545, 15 arti- 1862 Rhynchonella lucida Gould, p. 120. culated specimens, one ventral and one dorsal valves (MNHN IB- 2008 Basiliola lucida - Bitner, p. 427, fig. 5A-G cum( syn.). 2009-538, IB-2013-683); stn CP 1546, 2 articulated specimens and 2009 Basiliola lucida - Bitner, p. 7, fig. 3A-C. one dorsal valve (MNHN IB-2009-541); stn DW 1548, one articu- 2010 Basiliola lucida - Zezina, p. 1179. lated specimen (MNHN IB-2013-684); stn DW 1554, one articula- 2015 Basiliola lucida - Bitner, p. 36, fig. 2C-D. ted specimen (MNHN IB-2013-685); stn DW 1555, one articulated specimen (MNHN IB-2013-686); stn CH 1557, one articulated spe- Material examined: Tonga, Bordau 2 cruise, stn DW 1516, 4 articulated specimens (MNHN IB-2013-695); stn DW 1518, one cimen (MNHN IB-2013-687); stn DW 1569, one articulated speci- articulated specimen (MNHN IB-2009-554); stn DW 1523, one arti- men (MNHN IB-2013-688); stn CP 1593, one articulated specimen culated specimen (MNHN IB-2009-552); stn DW 1532, 19 articulated (MNHN IB-2013-689); stn DW 1597, one articulated specimen specimens, 3 ventral and one dorsal valves (MNHN IB-2009-558); (MNHN IB-2013-690); stn DW 1614, 2 articulated specimens (IB- stn CP 1533, 19 articulated specimens, one ventral and 2 dorsal valves 2009-543); stn DW 1615, one articulated specimen (IB-2009-539); (MNHN IB-2009-549); stn DW 1534, 2 articulated specimens (MNHN stn DW 1617, 4 articulated specimens (MNHN IB-2013-691); stn IB-2013-696); stn DW 1535, 2 articulated specimens (MNHN IB- DW 1618, one articulated specimen (MNHN IB-2013-692); stn DW 2009-557); stn DW 1536, 13 articulated specimens (MNHN IB-2013- 1619, 2 articulated specimens and 2 ventral valves (MNHN IB-2013- 697); stn DW 1537, 2 articulated specimens (MNHN IB-2009-553); 693-694). stn CP 1541, one articulated specimen (MNHN IB-2009-559); stn Depth range: 427-656 m. DW 1583, 4 articulated specimens (MNHN IB-2013-698); stn DW Measurements: Max. length 18.9 mm, width 17.1 mm, 1587, 9 articulated specimens (MNHN IB-2009-546, IB-2013-699); thickness, 12.5 mm. stn DW 1589, 5 articulated specimens (MNHN IB-2009-556); stn DW 1604, 7 articulated specimens (MNHN IB-2009-547); stn DW Remarks. Basiliola beecheri is relatively com- 1614, one articulated specimen (MNHN IB-2009-560); stn DW 1630, mon (more than 50 specimens) in the material col- one ventral valve (MNHN IB-2009-562); stn DW 1634, 11 articulated lected from the Tonga Islands. It is a medium-sized specimens (MNHN IB-2009-548); stn DW 1635, 10 articulated speci- mens (MNHN IB-2009-551, IB-2013-700); stn DW 1636, 6 articula- rhynchonellide characterized by a smooth except ted specimens and one dorsal valve (MNHN IB-2009-555). for concentric growth lines, dorsibiconvex, strongly Depth range: 227-549 m. uniplicate shell, and internally by wide hinge plates, Measurements: Max. length 13.9 mm, width 11.9 mm, thickness 9.3 mm. subfalciform crura and lack of cardinal process. Al- though widely distributed in the Pacific from New Remarks. Basiliola lucida is the second most Caledonia to Fiji and Hawaii (Dall 1895; Laurin common species (more than 120 specimens) in this 1997; Logan 2007; Bitner 2006b, 2008, 2009, 2015), collection. Its shell is smooth with only growth lines, this is the first record from Tonga. dorsibiconvex and strongly uniplicate. It can be eas- ily distinguished from B. beecheri by its small size, Genus Basiliolella d’Hondt, 1987 elongate outline and narrower outer hinge plates Type species - Basiliolella ferox d’Hondt, 1987 by original designation (Hatai 1940; Cooper 1959; Laurin 1997; Bitner 2008, of d’Hondt (1987: 39) 2009, 2015). Basiliola lucida is reported for the first time Basiliolella colurnus (Hedley, 1905) from the Tonga Islands although it was already iden- Fig. 2N-S tified in the nearby Fiji region (Bitner 2008). This 1905 Hemithyris colurnus Hedley, pp. 44-45, figs. 7-8. species was originally described from off Japan 1920 Hemithyris colurnus - Dall, p. 288. (Gould 1862) and also occurs in the New Caledo- 1959 Eohemithyris colurnus - Cooper, p. 32, pl. 15, figs. 15-26. 1981a Eohemithyris colurnus - Zezina, p. 11. nian region (Laurin 1997; Bitner 2009, 2015). 2010 Eohemithyris colurnus - Zezina, p. 1180.

Basiliola beecheri (Dall, 1895) Material examined: Tonga, Bordau 2 cruise, stn DW Fig. 2I-M 1516, 2 articulated specimens (MNHN IB-2013-701); stn DW 1532, Recent brachiopods from the Tonga Islands, SW Pacific 591

4 articulated specimens (MNHN IB-2013-702); stn DW 1534, 2 ar- Depth range: 483-531 m. ticulated specimens (MNHN IB-2009-550); stn DW 1536, 2 articu- lated specimens (MNHN IB-2013-703); stn DW 1543, 2 articulated Remarks. Although the material is very lim- specimens (MNHN IB-2013-704); stn CP 1545, 7 articulated spe- cimens and one ventral valve (MNHN IB-2013-705-706); stn DW ited and poorly preserved, a very large size (ob- 1589, one articulated specimen (MNHN IB-2013-707); stn 1604, served width 42.7 mm) and smooth marked by one articulated specimen (MNHN IB-2013-708); stn DW 1606, one concentric growth lines, thick shell with incurved articulated specimen (MNHN IB-2013-709); stn DW 1612, one arti- valve margins make Dyscolia johannisdavisi one of culated specimen (MNHN IB-2009-561); stn DW 1634, one articu- lated specimen (MNHN IB-2013-710). the most easily recognizable species among living Depth range: 227-447 m. brachiopods. This species was long considered to Measurements: Max. length 14.6 mm, width 15.2 mm, be restricted to the Indian Ocean (Helmcke 1940; thickness 8.5 mm. Muir-Wood 1959; Cooper 1983; Hiller 1986; Zezi- na 1994) until it was discovered in material from Remarks. Basiliolella colurnus is the third rhyn- chonellide brachiopod recognized in the material the New Caledonian region (Laurin 1997; Bitner under study. This is a medium-sized species with 2009). This is its second occurrence in the Pacific smooth with numerous distinct growth lines, nearly and the first from Tonga. equally convex shell subpentagonal in outline. The anterior commissure is uniplicate with a broad, Subfamily Aenigmathyridinae Cooper, 1983 Genus Thomson, 1927 gentle fold. The dorsal interior with a low median Abyssothyris Type species - Terebratula wyvilli Davidson, 1878 by original ridge and thickened crural bases. In size and outline designation (Thomson 1927: 170) B. colurnus is similar to B. beecheri but it differs in being nearly equally convex, weakly uniplicate and Abyssothyris wyvillei (Davidson, 1878) having thickened crural bases, whereas B. beecheri has unequal valves and strongly folded anterior 1878 Terebratula Wyvilli Davidson, p. 436. commissure. 2008 Abyssothyris wyvillei - Bitner, p. 429, fig. 6I-L cum( syn.). 2010 Abyssothyris wyvillei - Zezina, p. 1185. B. colurnus is known from off the eastern coast of Australia (Cooper 1959; Zezina 1981a; Material examined: Tonga, Bordau 2 cruise, stn CP 1625, Logan 2007). Zezina (2010) also reported this spe- one articulated specimen, strongly broken (MNHN IB-2013-712), stn DW 1630, one immature articulated specimen (MNHN IB- cies from the southern Sea of Japan, but without 2013-790). description or illustration. This is its first record Depth range: 360-824 m. from Tonga. Measurements: Max. length 8.2 mm, width 8.0 mm and thickness, 5.1 mm. Order Waagen, 1883 Terebratulida Remarks. This short-looped species is very Suborder Terebratulidina Waagen, 1883 rare and poorly preserved in the material from Superfamily Dyscolioidea Fischer & Œhlert, 1891 Tonga. It is characterized by a small, smooth, ex- Family Dyscoliidae Fischer & Œhlert, 1891 cept for growth lines, and thin shell with a deeply Subfamily Dyscoliinae Fischer & Œhlert, 1891 unisulcate anterior commissure. The lack of dor- Genus Dyscolia Fischer & Œhlert, 1890 sal median septum makes it readily distinguishable Type species - Terebratulina wyvilli Davidson, 1878 by original designation by Fischer & Œhlert (1890: 70) from the externally very similar genus Nipponithyris Yabe & Hatai, 1934 (see Bitner 2008). Dyscolia johannisdavisi (Alcock, 1894) Like most species described here, this is the first record of A. wyvillei from Tonga, but it is 1894 Terebratula johannisdavisi Alcock, p. 139. known from many localities in the Pacific Ocean 1940 Dyscolia johannis-davisi - Helmcke, p. 261, figs. 22, 25, 25b. and has a circumpolar distribution in the South 1959 Dyscolia johannisdavisi - Muir-Wood, pp. 300-302, pl. 1, figs. 1, 3, 4. Ocean (Cooper 1982, 1983; Foster 1989; Laurin 1986 Dyscolia cf. johannisdavisi - Hiller, pp. 106-110, figs. 4, 5. 1997; Bitner 2006b, 2008; Logan 2007; MacFarlan 1994 Dyscolia johannisdavisi - Zezina, pp. 46-48, fig. 2. 1997 Dyscolia johannisdavisi - Laurin, pp. 429-430, fig. 43J-L. et al. 2009). 2009 Dyscolia johannisdavisi - Bitner, p. 11, fig. 6E, F. Genus Xenobrochus Cooper, 1981 Material examined: Tonga Islands, Bordau 2 cruise, stn Type species - Gryphus africanus Cooper, 1973 by original DW 1617, one dorsal valve, broken (MNHN IB-2013-711). designation (Cooper 1981: 19) 592 Bitner M.A.

Xenobrochus rotundus Bitner, 2008 collection. This species differs from those described Fig. 3A-E below in its smaller size and coarsely ribbed orna- mentation. The ribs are also less numerous. First 2008 Xenobrochus rotundus Bitner, pp. 429-431, fig. 6A-H. described from off the Philippines (Dall 1920), T. 2010 Xenobrochus rotundus - Zezina, p. 1185. reevei is known also from the New Caledonian and Material examined: Tonga, Bordau 2 cruise, stn DW 1534, Fijian regions (Laurin 1997; Bitner 2006b, 2008, one immature articulated specimen (MNHN IB-2013-713); stn DW 2009). The specimens from off Celebes and Bor- 1537, 14 articulated specimens, 9 ventral and 2 dorsal valves (MNHN IB-2013-714); stn DW 1544, one articulated specimen (MNHN neo assigned to T. reevei by Zezina (1981a) are much IB-2013-715); stn DW 1552, one articulated specimen (MNHN IB- larger. This attribution remains unclear because nei- 2013-716). ther illustrations and nor detailed description were Depth range: 302-500 m provided. Measurements: Max. length 9.0 mm, width 6.5 mm, thickness 7.0. Terebratulina japonica (G.B. Sowerby, 1846) Remarks. Xenobrochus rotundus was originally Fig. 3H-K described from the Fiji and Wallis and Futuna Is- lands (Bitner 2008); Tonga is another occurrence of 1846 Terebratula japonica G.B. Sowerby, p. 91. this species. The shell is small, smooth with poorly 2008 Terebratulina japonica - Bitner, p. 433, fig. 7A-D cum( syn.). 2010 Terebratulina japonica - Zezina, p. 1185. marked growth lines, strongly biconvex with recti- 2017 Terebratulina japonica - Bitner & Romanin, p. 287, fig. 1A-B. marginate anterior commissure. Its beak is suberect 2018 Terebratulina japonica - Bitner & Romanin, p. 556, fig. 6C-D. with a small, circular foramen. The deltidial plates Material examined: Tonga, Bordau 2 cruise, stn DW 1509, form a wholly visible symphytium. The cardinalia one articulated specimen (MNHN IB-2009-537); stn DW 1516, 5 include a distinct, semi-elliptical cardinal process articulated specimens (MNHN IB-2013-725-726); stn DW 1523, 7 and very narrow hinge plates. The loop is short, and articulated specimens (MNHN IB-2013-727); stn DW 1524, 3 arti- its transverse band has a weak fold. The specimens culated specimens (MNHN IB-2013-728); stn DW 1548, 2 articula- ted specimens (MNHN IB-2013-729); stn DW 1554, one articulated from Tonga differ from those from Fiji in being specimen (MNHN IB-2013-730); stn DW 1555, 3 articulated speci- more elongate. mens (IB-2013-731); stn CH 1557, 14 articulated specimens (MNHN IB-2013-732); stn DW 1569, one articulated specimen (MNHN IB- 2013-733); stn DW 1570, one articulated specimen (MNHN IB- 2013-734); stn CP 1582, one articulated specimen (MNHN IB-2013- Superfamily Cancellothyridoidea Thomson, 1926 735); stn DW 1584, 2 articulated specimens (MNHN IB-2013-736); Family Cancellothyrididae Thomson, 1926 stn DW 1585, one articulated specimen (MNHN IB-2013-737); stn Subfamily Cancellothyridinae Thomson, 1926 DW 1597, one articulated specimen (MNHN IB-2013-738) ; stn DW 1604, one articulated specimen (MNHN IB-2013-739); stn DW 1615, Genus Terebratulina d’Orbigny, 1847 one articulated specimen (MNHN IB-2013-740); stn DW 1616, one Type species - Anomia retusa Linnaeus, 1758, by subsequent articulated specimen (MNHN IB-2013-741); stn DW 1617, one ar- designation of Brunton et al. (1967: 176) ticulated specimen (MNHN IB-2013-742); stn DW 1618, 3 articu- lated specimens (MNHN IB-2013-743); stn DW 1619, 6 articulated Terebratulina reevei Dall, 1920 specimens and one ventral valve (MNHN IB-2013-744-745); stn CP 1620, one articulated specimen (MNHN IB-2013-746); stn CH 1621, Fig. 3F-G 5 articulated specimens (MNHN IB-2009-536); stn CP 1640, one ar- ticulated specimen (MNHN IB-2013-747). 1920 Terebratulina reevei Dall, pp. 305-306. Depth range. 79-781 m. 2008 Terebratulina reevei - Bitner, p. 434, fig. 8A-G cum( syn.). Max. length 13.5 mm, width 10.9 mm, 2009 Terebratulina reevei - Bitner, p. 13, fig. 7B. Measurements: 2010 Terebratulina reevei - Zezina, p. 1186. thickness 6.3 mm.

Material examined: Tonga, Bordau 2 cruise, stn DW 1516, Remarks. The second Terebratulina species in 2 articulated specimens (MNHN IB-2013-749); stn DW 1523, 3 arti- culated specimens (MNHN IB-2013-750); stn DW 1567, 13 articula- the material under study is T. japonica. It is relatively ted specimens and one ventral valve (MNHN IB-2013-751); stn DW common, with more than 60 specimens. This spe- 1632, one articulated specimen (MNHN IB-2013-752). cies can be easily differentiated from T. reevei by its Depth range: 229-618 m. ornamentation of numerous, fine ribs and from T. Measurements: Max. length 8.8 mm, width 6.2 mm, thickness 3.6 mm. australis by its elongate oval outline and large fora- men. The anterior commissure in adult T. japonica is Remarks. Terebratulina reevei is one of the weakly and broadly uniplicate. Originally described three Terebratulina species recognized in the Tonga from off Japan (G.B. Sowerby 1846; Hatai 1940; Recent brachiopods from the Tonga Islands, SW Pacific 593

Fig. 3 - A-E - Xenobrochus rotundus Bitner, 2008, Tonga, Bordau 2, SEM, A-B dorsal view of articulated specimen, and enlargement of posterior part to show details of the beak, MNHN IB-2013-714, stn DW 1537, 391-421 m; C-E inner views of disarticulated specimen, MNHN IB-2013-716, stn DW 1552, 491-500 m, (C) ventral valve, (D-E) dorsal valve, and enlargement (E) of posterior part to show details of brachial skeleton. F-G - Terebratulina reevei Dall, 1920, Bordau 2, MNHN IB-2013-750, SEM, stn DW 1523, 300-302 m, dorsal view of articulated specimen, and enlargement (G) of posterior part. H-K - Terebratulina japonica (G.B. Sowerby, 1846), Bordau 2, H-J dorsal, lateral and anterior views of articulated specimen, MNHN IB-2013-725, stn DW 1516, 229-246 m; K inner view of dorsal valve, MNHN IB-2013-745, SEM, stn DW 1619, 591-593 m. L-S - Terebratulina australis Bitner, 2006, Bordau 2, L dorsal view of young speci- men, MNHN IB-2013-748, SEM, stn DW 1631, 407-443 m; M-O dorsal, lateral and anterior views of articulated specimen, MNHN IB-2013-722, stn DW 1628, 400-416 m; P dorsal view of articulated specimen, MNHN IB-2013-724, stn DW 1634, 321-322 m; Q-R inner and tilted views of dorsal valve, MNHN IB-2013-718, SEM, stn CP 1510, 461-497 m. 594 Bitner M.A.

(MNHN IB-2009-529, IB-2013-748); stn DW 1632, one articulated specimen (MNHN IB-2013-723); stn DW 1634, 42 articulated speci- mens and one ventral valve (MNHN IB-2009-519, IB-2013-724); stn DW 1635, 42 articulated specimens and one dorsal valve (MNHN IB-2009-517); stn DW 1636, 20 articulated specimens (MNHN IB- 2009-518). Depth range: 300-618 m. Measurements: Max length 12.4 mm, width 11.7 mm, thickness 6.1 mm (see also Fig. 4).

Remarks. Terebratulina australis is the most common species (more than 700 specimens) in this collection. It was originally described from off the Fiji Islands (Bitner 2006b, 2008) where it is rare. The shell is thickened posteriorly. The inner sock- et ridges are narrow but massive, the ring is broad and subsquare. By its rounded outline, shell surface covered with numerous, fine but distinct ribs and a very small foramen this species differs from other Terebratulina species. This species, although not described yet, was Fig. 4 - Intraspecific variation in Terebratulina australis Bitner, 2006. Scatter diagram plotting length to width and thickness. N also recognized in the material from New Zealand number of specimens. (J.H. Robinson, personal communication).

Logan 2007) it was also found in the South China Suborder Terebratellidina Muir-Wood, 1955 Sea (Bitner & Romanin 2017, 2018) and Fiji (Bitner Superfamily Laqueoidea Thomson, 1927 2006b, 2008). Family Frenulinidae Hatai, 1938 Subfamily Frenulininae Hatai, 1938 Terebratulina australis Bitner, 2006 Genus Frenulina Dall, 1895 Figs. 3L-R, 4 Type species - Anomia sanguinolenta Gmelin, 1791 by original designation of Dall (1895: 724) 2006b Terebratulina australis Bitner, pp. 25-27, fig. 5D-J. 2008 Terebratulina australis - Bitner, p. 434, fig. 7E-I. Frenulina sanguinolenta (Gmelin, 1791) 2010 Terebratulina australis - Zezina, p. 1186. Fig. 5A-B

Material examined: Tonga, Bordau 2 cruise, stn CP 1510, 3 articulated specimens (MNHN IB-2013-717-718); stn DW 1517, 1791 Anomia sanguinolenta Gmelin, p. 3347. one articulated specimen (MNHN IB-2009-534); stn DW 1518, 4 2014 Frenulina sanguinolenta - Bitner, pp. 250, 252-253, 255, fig. 8A-E articulated specimens (MNHN IB-2013-719); stn DW 1520, one arti- (cum syn.). 2015 Frenulina sanguinolenta - Bitner, p. 41, fig. 4F-G. culated specimen (MNHN IB-2013-720); stn DW 1523, 5 articulated 2016 Frenulina sanguinolenta - Bitner & Logan, p. 24, fig. 13A. specimens (MNHN IB-2009-525); stn DW 1524, 3 articulated speci- 2016 Frenulina sanguinolenta - Álvarez, pp. 63-65, pls. 25K-BB, 26A-O. mens (MNHN IB-2009-533); stn CP 1525, 3 articulated specimens 2018 Frenulina sanguinolenta - Bitner & Romanin, pp. 558-559, fig. 6O- (MNHN IB-2009-524); stn DW 1532, 90 articulated specimens, 25 P. ventral and 20 dorsal valves (MNHN IB-2009-528); stn CP 1533, 2 articulated specimens and 2 dorsal valves (MNHN IB-2009-526); Material examined: Tonga, Bordau 2 cruise, Stn DW 1569, stn DW 1534, 12 articulated specimens (MNHN IB-2009-523); stn one articulated specimen (MNHN IB-2013-756). DW 1536, 35 articulated specimens and one ventral valve (MNHN Measurements: Length 6.3 mm, width 6.1 mm, thickness IB-2009-530); stn DW 1537, 49 articulated specimens, one ventral 3.6 mm. and one dorsal valves (MNHN IB-2009-521); stn DW 1540, 13 ar- Depth range: 433 m. ticulated specimens (MNHN IB-2009-535); stn CP 1541, 2 articula- ted specimens (MNHN IB-2009-531); stn DW 1606, 4 articulated specimens (MNHN IB-2013-721); stn DW 1611, 2 articulated spe- Remarks. This species, although already re- cimens (MNHN IB-2009-522); stn DW 1612, 8 articulated speci- ported from the Tonga region (Thomson 1927; Saito mens (MNHN IB-2009-532); stn DW 1614, 3 articulated specimens & Endo 2001; Logan 2007; Zezina 2010), is repre- (MNHN IB-2009-527); stn DW 1628, 147 articulated specimens, 3 ventral and 3 dorsal valves (MNHN IB-2009-520, IB-2013-722); sented by only one specimen in the studied material. stn DW 1630, 55 articulated specimens (MNHN IB-2009-516); stn By its red colour pattern and a small, sulcate shell DW 1631, 99 articulated specimens, 2 ventral and 12 dorsal valves Frenulina sanguinolenta is easily distinguishable among Recent brachiopods from the Tonga Islands, SW Pacific 595

Fig. 5 - A-B - Frenulina sanguinolenta (Gmelin, 1791), Bordau 2, MNHN IB-2013-756, stn DW 1569, 433 m, dorsal view of articulated specimen, and enlargement (B) of posterior part, SEM. C-G - Septicollarina zezinae Bitner, 2009, Tonga, Bordau 2, MNHN IB-2013-755, stn DW 1523, 300-302 m, SEM; (C-D) inner and oblique views of dorsal valve; (E) dorsal view of articulated specimen; (F-G) inner and tilted views of ventral valve to show dental plates and pedicle collar supported by a septum. H-J - Fallax neocaledonensis Laurin, 1997, MNHN IB-2013-753, Bordau 2, stn DW 1617, 483-531 m; (H-I) dorsal and anterior views of articulated specimen; (J) inner view of dorsal valve of the same disarticulated specimen. brachiopods. It is one of the most widely distributed Genus Septicollarina Zezina, 1981 species in the West Pacific, known from Japan, Aus- Type species - Septicollarina hemiechinata Zezina, 1981 by original tralia and New Caledonia to French Polynesia and designation of Zezina (1981a: 16) Hawaii (Hatai 1940; Emig 1987; Saito 1996; Laurin 1997; Logan 2007; Bitner 2006a, 2006b, 2008, 2009, Septicollarina zezinae Bitner, 2009 2010, 2014, 2015; Simon & Hoffmann 2013; Bitner Fig. 5C-G & Romanin 2018; Álvarez 2016; Simon et al. 2016, 2008 Septicollarina sp. - Bitner, pp. 437-439, fig. 16A. 2018). Recently it has also been recognized in the 2009 Septicollarina zezinae Bitner, pp. 14, 24-25, fig. 8A-N. Madagascar area (Bitner & Logan 2016). 2010 Septicollarina sp. - Zezina, p. 1189. 2014 Septicollarina zezinae - Bitner, p. 250, figs. 6A-I, 7A-H. 2015 Septicollarina zezinae - Bitner, p. 41, fig. 4H-I.

Material examined: Tonga, Bordau 2 cruise, stn DW 1516, Superfamily Kingenoidea Elliott, 1948 one articulated specimen (MNHN IB-2013-754); stn DW 1523, 4 ar- Family Aulacothyropsidae Dagys, 1972 ticulated specimens (MNHN IB-2013-755). Subfamily Babukellinae MacKinnon, Smirnova & Depth range: 229-302 m. Measurements: Max. length 4.6 mm, width 5.9 mm, Lee, 2006 thickness 3.4 mm. 596 Bitner M.A.

Remarks. This species is rare in the Tonga Amphithyris buckmani Thomson, 1918 material, and is represented only by immature spe- Fig. 6A-F cimens. Its shell is transversely subpentagonal with the surface covered with delicate radial lines and 1918 Amphithyris buckmani Thomson, p. 22, pl. 15, fig. 9, pl. 16, fig. 35. rare pustules. The ventral valve interior has distinct 2006b Amphithyris buckmani - Bitner, pp. 28-30, fig. 6a-f. dental plates and a wide pedicle collar supported by 2008 Amphithyris buckmani - Bitner, pp. 440-442, fig. 11A-L. 2008 Amphithyris buckmani - MacKinnon et al., p. 329, fig. 1A-D. a short median septum. The cardinalia are without 2010 Amphithyris buckmani - Zezina, p. 1192. a cardinal process and with inner hinge plates atta- 2014 Amphithyris buckmani - Nauendorf et al., pp. 224-225. ched to the median septum, forming a septalium. Material examined: Tonga, Bordau 2 cruise, stn CP 1560, The loop has broad ascending branches and tran- 5 articulated specimens (MNHN IB-2013-757); stn CP 1562, 32 arti- sverse band and is weakly spinose anteriorly. culated specimens and one dorsal valve (MNHN IB-2013-758-759); Although widely distributed in the SW Paci- stn CP 1578, 2 articulated specimens (MNHN IB-2013-760); stn DW fic, occurring from New Caledonia, Fiji to French 1612, 2 articulated specimens (MNHN IB-2013-761); stn DW 1628, 5 articulated specimens (MNHN IB-2013-762); stn DW 1630, one Polynesia (Bitner 2008, 2009, 2014, 2015), this is the articulated specimen (MNHN IB-2013-763); stn DW 1631, 6 articu- first record ofS. zezinae from Tonga. lated specimens (MNHN IB-2013-764). Depth range: 327-443 m. Genus Fallax Atkins, 1960 Measurements: Max. length 4.0 mm, width 4.3 mm, thickness 1.3 mm. Type species - Fallax dalliniformis Atkins, 1960, by original designation (Atkins 1960: 72) Remarks. With more than 50 specimens Amphithyris buckmani is a relatively common spe- Fallax neocaledonensis Laurin, 1997 Fig. 5H-J cies in the Tonga material but this is its first record from the area. Originally this species was described 1997 Fallax neocaledonensis Laurin, pp. 444-448, figs. 31-34, 46a-o. from the waters of New Zealand (Thomson 1918; 2006b Fallax neocaledonensis - Bitner, p. 27, fig. 5a-c. MacKinnon et al. 2008; Nauendorf et al. 2014), lat- 2008 Fallax neocaledonensis - Bitner, p. 437, fig. 10H-J. 2009 Fallax neocaledonensis - Bitner, p. 13, fig. 7I-L. er being also found in the Fiji region where it is the 2015 Fallax neocaledonensis - Bitner, p. 41, fig. 4J-K. commonest species (Bitner 2006b, 2008). This micromorphic species has a convex ven- Material examined: Tonga, Bordau 2 cruise, stn DW 1554, one articulated specimen (MNHN IB-2013-789); stn DW 1617, one tral valve with radial lines and a smooth, flat dorsal articulated specimen (MNHN IB-2013-753). valve. Its foramen is very large, amphithyrid, subcir- Depth range: 482-531 m. cular to oval in outline. The internal morphology is Measurements: Max. length 21.6 mm, width 23.0 mm, thickness 15.9 mm. very simple with a few characters only. The socket ridges are short, projecting slightly beyond the mar- Remarks. Fallax neocaledonensis is characteri- gin. The median septum is short and low. Crura and zed by a smooth, with numerous growth lines shell, loop are not developed. The specimens from Tonga widely triangular in outline and the presence of are smaller than those described from New Zealand well-developed dental plates. Its loop is of diplo- and Fiji where A. buckmani can reach more than 5 form type with broad ascending branches. In the mm in length (Bitner 2008; MacKinnon et al. 2008). material under study this species is very rare. In the New Caledonia region from where it was originally Genus Annuloplatidia Zezina, 1981 described, F. neocaledonensis is one of the most com- Type species - Annuloplatidia indopacifica Zezina, 1981 by original mon species (Laurin 1997; Bitner 2009, 2015). It designation of Zezina (1981b: 144) was also identified in the material from Fiji (Bitner Annuloplatidia curiosa Bitner, 2014 2006b, 2008). Fig. 6G-M

Superfamily Platidioidea Thomson, 1927 2014 Annuloplatidia curiosa Bitner, pp. 255-258, figs. 9A-H, 10A-F cum( Family Platidiidae Thomson, 1927 syn.). Subfamily Platidiinae Thomson, 1927 2015 Annuloplatidia curiosa - Bitner, p. 42, fig. 5K-L. 2018 Annuloplatidia curiosa - Bitner & Romanin, pp. 559-561, fig. 7H-M. Genus Amphithyris Thomson, 1918 Type species - Amphithyris buckmani Thomson, 1918 by original Material examined: Tonga, Bordau 2 cruise, stn DW 1521 designation of Thomson (1918: 20) 2 articulated specimens (MNHN IB-2013-765); stn DW 1605, 7 ar- Recent brachiopods from the Tonga Islands, SW Pacific 597

Fig. 6 - A-F - Amphithyris buckmani Thomson, 1918, Tonga, Bordau 2; (A) dorsal view of young articulated specimen, MNHN IB-2013-759, stn CP 1562, 417-424 m; (B) ventral view of articulated specimen, MNHN IB-2013-760, stn CP 1578, 329-331 m; (C-D) dorsal view of articulated specimen, and enlargement (D) of posterior part to show details of the beak, MNHN IB-2013-759, stn CP 1562; (E-F) inner and oblique views of dorsal valve, MNHN IB-2013-759, stn CP 1562. G-M - Annuloplatidia curiosa Bitner, 2014, Bordau 2;( G-H) ventral views of two articulated specimens, MNHN IB-2013-766, stn DW 1605, 441 m; (I) dorsal view of complete specimen, MNHN IB-2013-767, stn DW 1615, 482-504 m; (J-M) inner views of disarticulated specimen, MNHN IB-2013-768, stn DW 1616, 664-781 m, (J) ventral valve, (K-M) inner, tilted and oblique views of dorsal valve. N-O - Leptothyrella fijiensis Bitner, 2008, dorsal view of articulated specimen, and enlargement (O) of posterior part, MNHN IB-2013-769, stn DW 1566, 530-531 m. All SEM. ticulated specimens (MNHN IB-2013-766); stn DW 1615, 7 articu- Remarks. Annuloplatidia curiosa is relatively lated specimens (MNHN IB-2013-767); stn DW 1616, 11 articulated rare in the investigated material; it was found in specimens (MNHN IB-2013-768). Depth range: 225-781 m. only 4 stations. This species is large for the genus, Measurements: Max. length 5.2 mm, width 7.4 mm, being readily distinguishable from other Annulopla- thickness 2.2 mm. tidia species by ribbed ornamentation of the ventral 598 Bitner M.A.

valve. Its dorsal valve surface is rough and irregu- 2008 Dallina triangularis - Bitner, pp. 444-446, fig. 16B-G. 2010 Dallina triangularis - Zezina, p. 1195. lar with poorly marked growth lines. Internally it is 2018 Dallina triangularis - Bitner & Romanin, p. 559, fig. 7A-B. characterized by a loop with descending branches attached to ascending branches that diverge from Material examined: Tonga, Bordau 2 cruise, stn DW 1607, the septum and the ascending branches are con- one articulated specimen, partly broken (MNHN IB-2009-1198). Depth range: 356-367 m. nected by a transverse band. This species is widely distributed in the SW Remarks. This species is very rare in the in- Pacific, being recorded from New Caledonia, New vestigated material, represented by one partly bro- Zealand, Wallis and Futuna Islands and French ken specimen. It is characterized by a medium-sized Polynesia (Bitner 2007, 2008, 2014, 2015). Recently (length 32.5 mm), smooth, ornamented only by it was also identified in the material from the South weakly defined growth lines shell with a paraplicate China Sea, constituting the first occurrence in the anterior commissure and a large, circular perme- northern hemisphere waters (Bitner & Romanin sothyrid foramen. Internally the characteristic fea- 2018). tures are a lack of dental plates and a long loop not attached to the septum. Subfamily Phaneroporinae Zezina, 1981 Dallina triangularis was established based on Genus Leptothyrella Muir-Wood, 1965 the material from off Japan (Yabe & Hatai 1934; Type species - Leptothyris ignota Muir-Wood, 1959 by original Hatai 1940). It is also known from Fiji (Bitner 2008) designation of Muir-Wood (1959: 308) and recently recognized in the South China Sea (Bit- ner & Romanin 2018). Leptothyrella fijiensis Bitner, 2008 Fig. 6N-O Subfamily Nipponithyridinae Hatai, 1938 2008 Leptothyrella fijiensis Bitner, p. 442-444, fig. 13A-L. Genus Campages Hedley, 1905 2010 Leptothyrella fijiensis- Zezina, p. 1193. Type species - Campages furcifera Hedley, 1905 by original designation of Hedley (1905: 43) Material examined: Tonga, Bordau 2 cruise, stn CP 1566, 2 articulated specimens (MNHN IB-2013-769). Depth range: 530-531 m. Campages ovalis Bitner, 2008 Measurements: Max. length 4.0 mm, width 3.6 mm, Fig. 7F-M thickness 1.2 mm. 2008 Campages ovalis Bitner, pp. 449-451, fig. 18A-J. Remarks. This species is very rare in the col- 2010 Campages ovalis - Zezina, p. 1196. lection from Tonga, found in only one station. Pre- Material examined: Tonga, Bordau 2 cruise, stn DW viously L. fijiensis has been known only from Fiji, its 1517, one articulated specimen (MNHN IB-2009-1240); stn DW type locality (Bitner 2008). Its shell is small, smo- 1523, 3 articulated specimens (MNHN IB-2013-770); stn CP 1525, one articulated specimen and one ventral valve (MNHN IB-2013- oth with poorly defined concentric growth lines, 771); stn DW 1532, 14 articulated specimens, 2 ventral and 2 dorsal weakly biconvex with a large, triangular hypothyrid valves (MNHN IB-2013-772); stn DW 1535, one articulated speci- foramen. The beak ridges are sharp with two rows men and one ventral valve (MNHN IB-2013-773); stn DW 1536, of small tubercles. The specimens from Tonga are 4 articulated specimens (MNHN IB-2013-774); stn DW 1537, 3 articulated specimens (MNHN IB-2013-775); stn DW 1540, one more rounded than those from Fiji. articulated specimen (MNHN IB-2013-776); stn DW 1583, 4 arti- culated specimens (MNHN IB-2013-777); stn DW 1587, 2 articula- Superfamily Terebratelloidea King, 1850 ted specimens (MNHN IB-2013-778); stn DW 1604, one articula- Family Dallinidae Beecher, 1893 ted specimen (MNHN IB-2013-779); stn DW 1606, one articulated specimen (MNHN IB-2013-780); stn DW 1612, one articulated Subfamily Dallininae Beecher, 1893 specimen (MNHN IB-2013-781); stn DW 1628, 4 articulated spe- Genus Dallina Beecher, 1893 cimens (MNHN IB-2013-782); stn DW 1634, 5 articulated speci- Type species - Terebratula septigera Lovén, 1846 by original mens (MNHN IB-2013-783); stn DW 1635, 2 articulated speci- designation of Beecher (1893: 383) mens (MNHN IB-2013-784); stn DW 1636, 5 articulated specimens (MNHN IB-2013-785). Dallina triangularis Yabe & Hatai, 1934 Depth range: 227-421 m. Measurements: Max. length 10.4 mm, width 8.2 mm, Fig. 7N thickness 8.0 mm. 1934 Dallina triangularis Yabe & Hatai, p. 662, figs. 31-35. 1940 Dallina triangularis - Hatai, p. 320, pl. 7, figs. 22-27. Remarks. This species is moderately com- Recent brachiopods from the Tonga Islands, SW Pacific 599

Fig. 7 - A-B - Minutella minuta (Cooper, 1981), Tonga, dorsal view of articulated specimen, and enlargement (B) of posterior part to show rugideltidium, MNHN IB-2013-788, Bordau 2, stn DW 1521, 225-233 m, SEM. C-E - Thecidellina maxilla (Hedley, 1899), Bordau 2, MNHN IB-2013-787, stn DW 1567, 351-356 m, SEM; (C) dorsal view of articulated specimen; (D-E) inner and posterior views of dorsal valve. F-M - Campages ovalis Bitner, 2008, MNHN IB-2013-772, Bordau 2, stn DW 1532, 322 m, SEM; (F-I) dorsal views of two articulated specimens, and enlargement (G, I) of posterior part; (J-K) inner view of ventral valve and enlargement of posterior part to show delthyrial cavity; (L-M) inner and oblique views of dorsal valve. N - Dallina triangularis Yabe & Hatai, 1934, dorsal view of articulated specimen, partly broken, MNHN IB-2009-1198, Bordau 2, stn DW 1607, 356-367 m. mon (nearly 60 specimens) and widely distributed 2008). C. ovalis is a small species with a rounded to (found in 17 stations). Hitherto it has been known oval, strongly biconvex shell. It has a small, circu- only from Fiji and Wallis and Futuna Islands (Bitner lar foramen and a visible symphytium. Its teeth are 600 Bitner M.A.

small and without dental plates. The loop is typi- because of insufficient description and illustration. cal for the genus with narrow, parallel descending It is worth mentioning that Simon et al. (2018) did branches and a broad hood. The muscle scars are not express their opinion about specimens of T. strongly defined. maxilla well illustrated and described by Lee & Rob- inson (2003) and Bitner (2008, 2010). Order Thecideida Elliott, 1958 Nevertheless, all researchers (Lee & Robin- Superfamily Thecideoidea Gray, 1840 son 2003; Simon & Hoffmann 2013; Simon et al. Family Thecidellinidae Elliott, 1958 2018) suggest that to elucidate this taxonomic prob- Subfamily Thecidellininae Elliott, 1953 lem morphological and ontogenetic studies of large Genus Thecidellina Thomson, 1915 populations together with molecular genetic analy- Type species - Thecidium barretti Davidson, 1864, by original ses are strongly needed. designation of Thomson (1915: 462) Subfamily Minutellinae Logan & Baker, 2013 Thecidellina maxilla (Hedley, 1899) Fig. 7C-E Genus Minutella Hoffmann & Lüter, 2010 Type species: Minutella tristani Hoffmann & Lüter, 2010 by original designation of Hoffmann & Lüter (2010: 141) 1899 Thecidea maxilla Hedley, pp. 508-510, fig. 57. 2008 Thecidellina maxilla - Bitner, p. 451, fig. 19A-C cum( syn.). 2009 Thecidellina maxilla - Bitner, pp. 17-18, fig. 12A-J. Minutella minuta (Cooper, 1981) 2010 Thecidellina maxilla - Bitner, pp. 651-653, fig. 5G-J Fig. 7A-B 2010 Thecidellina maxilla - Zezina, p. 1181. 2014 Thecidellina maxilla - Bitner, p. 259, fig. 11A-I. 1981 Thecidellina minuta Cooper, p. 61, pl. 6, fig. 27-40. Material examined: Tonga, Bordau 2 cruise, stn DW 1514, 2009 Thecidellina minuta - Bitner, p. 18, fig. 13A-I. one articulated specimen (MNHN IB-2013-786); stn DW 1567, one 2010 Thecidellina minuta - Bitner, p. 653, fig. 5A-F. articulated specimen and one dorsal valve (MNHN IB-2013-787). 2010 Minutella minuta - Hoffmann & Lüter, pp. 148, 150, pl. 2, figs. 13-18, pl. 3, figs. 13-15. 130-356 m. Depth range: 2010 Minutella cf. minuta - Hoffmann & Lüter, pp. 150, 152, pl. 3, Measurements: Max. length 4.1 mm, width 3.2 mm, figs. 16-24. thickness 2.0 mm. 2013 Minutella minuta - Logan & Baker, p. 438, figs. 1F, 4I. 2013 Minutella minuta - Logan & Bitner, pp. 163-166, fig. 2A-O. Remarks. Thecidellina maxilla, like all species 2013 Minutella cf. minuta - Simon & Hoffmann, pp. 405-412, pl. 1, figs. 1-5, pl. 2, figs. 1-8. attributed to this genus, is characterized by a flat in- 2018 Minutella cf. minuta - Simon et al., pp. 495-496, pl. 7, figs. 1-2. terarea without pseudodeltidium, named planodelti- dium by Logan & Baker (2013), and a single, straight Material examined: Tonga, Bordau 2 cruise, stn DW 1521, dorsal septum (Hedley 1899; Cooper 1954; Lee & one articulated specimen (MNHN IB-2013-788). Robinson 2003). It is well-known in the South-West Depth range: 225-233 m. Measurements: Length 1.8 mm, width 1.5 mm, thickness Pacific although determination and validity of the 0.9 mm. Pacific species of Thecidellina has been the subject of discussion (e.g. Lee & Robinson 2003; Hoff- Remarks. Minutella minuta is very rare in the mann et al. 2009; Simon et al. 2018). In the opinion material collected in the Tonga area. This species of Lee & Robinson (2003) morphological charac- is characterized by a very small size and interarea ters on which particular species of Thecidellina have with convex, triangular rugideltidium sensu Logan been defined are minor and all Indo-Pacific forms & Baker (2013). The present finding extends its could be variants of Thecidellina maxilla. In contrast geographical range eastward. other researchers (Hoffmann et al. 2009; Simon & Recently Minutella is recorded from many Hoffmann 2013; Simon et al. 2018) find the -dif localities in the Pacific, from Okinawa, Indonesia, ferences in shell morphology to be noticeable. In Australia and New Caledonia to Palau, Fiji and their recent paper on thecideides Simon et al. (2018) Tonga (Hoffmann & Lüter 2010; Logan & Bit- discussed the taxonomic position of T. maxilla and ner 2013; Simon & Hoffmann 2013; Simon et al. concluded that several specimens attributed to the 2016, 2018). Hoffmann & Lüter (2010), Simon & latter species by d’Hondt (1987), Laurin (1997), and Hoffmann (2013) and Simon et al. (2018) stated Bitner (2007, 2009, 2014, 2015) represent either the that while all Pacific representatives ofMinutella are species T. congregata Cooper, 1954 or a new species, nearly identical to the specimens of M. minuta de- or in some cases the specimens cannot be evaluated scribed from Samper Bank, south-east of Madagas- Recent brachiopods from the Tonga Islands, SW Pacific 601

car, western Indian Ocean (the type locality of this tribution like Leptothyrella fijiensis known so far only species [Cooper 1981]) they prefer to describe them from Fiji (Bitner 2008). Also Xenobrochus rotundus as M. cf. minuta because a few minor differences are and Campages ovalis have the restricted distribution, present, although they do indicate that those few being reported earlier from Fiji and Wallis and Fu- minor morphological differences can be affected by tuna Islands (Bitner 2008). This collection also ex- species variability. Logan & Bitner (2013), however, tends the known ranges of several forms, including consider those minor differences as insufficient to Neoancistrocrania norfolki and Basiliolella colurnus. The separate distinct species and suggest that the Indo- latter species has been known so far from the east- Pacific forms are only varieties of M. minuta. This ern coast of Australia (Logan 2007). Interestingly, opinion is followed here. two species, Terebratulina japonica and Dallina trian- As mentioned above morphological stud- gularis have a similar distribution; both species were ies on large populations, combined with molecular described originally from off Japan (Hatai 1940) analysis could resolve such taxonomic problems. and have been also recognized in the South China Sea, Fiji and Tonga (Bitner 2008; Bitner & Romanin 2018; this paper). Although low affinity is observed with the Discussion fauna from New Zealand, sharing 6 species (Mac- Farlan et al. 2009; Bitner 2014), two species, Terebra- Twenty brachiopod species belonging to 17 tulina australis and Amphithyris buckmani occur only genera have been identified in the material collected in New Zealand, Fiji and Tonga. T. australis was during the French cruise Bordau 2 to the Tonga Is- originally described from Fiji where it is rare, but in lands, SW Pacific (Fig. 1). With 14 species, terebrat- the Tonga collection it is predominant, constituting ulide brachiopods dominate in the collection, while nearly 60% of the material. Similarly, A. buckmani craniids have one representative, rhynchonellides was originally described based on one specimen have three, and thecideide brachiopods have two from the New Zealand waters by Thomson (1918) representatives. Together with Novocrania turbinata and for 90 years it had been considered as endemic the total number of species documented in this re- to New Zealand until it was discovered in the Fiji gion is 21. This is only a slightly lower diversity than region where it is the most common species (Bitner that in the Fiji region (Bitner 2008; Hoffmann & 2008). The finding in Tonga extends the geographi- Lüter 2010). If compared with the New Caledonian cal range of both species. (Bitner et al. 2008; Bitner 2009, 2010, 2011, 2015; Bitner & Cohen 2015) and New Zealand regions Acknowledgements: My sincere thanks are to Philippe Bouchet for the opportunity to study the material and to Pierre Lozouet and (MacKinnon et al. 2008; MacFarlan et al. 2009; Jérôme Mainguy (all Muséum national d’Histoire naturelle, Paris) for Nauendorf et al. 2014; Robinson et al. 2016) the di- providing facilities during the visit at the Museum. Jeffrey H. Robin- versity of brachiopods in Tonga is half as great, be- son (University of Otago, Dunedin) is thanked for the helpful discus- ing however significantly higher than that in French sion on craniid brachiopods. The macrophotographs were taken by Grażyna Dziewińska (Institute of Paleobiology, Warszawa) to whom Polynesia (Bitner 2006a, 2007, 2014). Thus, the di- I am grateful. I thank the Editor, Gaia Crippa and two reviewers, Fer- versity of brachiopods from Tonga fits well to the nando Álvarez (University of Oviedo, Oviedo) and J.H. Robinson for pattern of decrease in species number, observed in their helpful suggestions. J. Robinson also improved English. the Pacific from west to east (see also discussion in Bitner 2014). References Apart from Frenulina sanguinolenta the remain- ing species represent the first records for Tonga. Alcock A. (1894) - A new brachiopod. In: Natural history The studied brachiopod fauna shows the greatest notes from the Royal Indian Marine Survey Steamer In- affinity to that from Fiji and New Caledonia, having vestigator, series 2, no.3. J. Asiatic Soc. Bengal, 63: 139-140. 16 and 12 species in common, respectively (Laurin Álvarez F. (2016) - Recent Brachiopods in the Œhlert 1997; Bitner 2008, 2009, 2010, 2011, 2015). Most Collection. In: Álvarez F., Emig C.C. & Tréguier J. - Bra- chiopodes actuels: historique et révision de la collection species have a wide distribution, being known from D.-P. Œhlert (Laval); brachiopodes des côtes françaises several localities in the Indo-Pacific province, how- métropolitaines. Bull. Soc. sci. nat. Ouest Fr., hors-série ever, there are also some with a very restricted dis- 2016-1: 25-109 and 145-167. 602 Bitner M.A.

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APPENDIX - Station list

Station Location Depth Species

Tonga Islands

BORDAU 2 Basiliola beecheri DW 1509 21°05'S, 175°22'W 456-510 m Terebratulina japonica Basiliola beecheri CP 1510 21°05'S, 175°23'W 461-497 m Terebratulina australis DW 1514 21°18'S, 175°05'W 130-133 m Thecidellina maxilla Basiliola lucida Basiliolella colurnus DW 1516 21°21'S, 175°02'W 229-246 m Terebratulina japonica Terebratulina reevei Septicollarina zezinae Terebratulina australis DW 1517 21°21'S, 175°07'W 342 m Campages ovalis Basiliola lucida DW 1518 21°21'S, 175°07'W 336-347 m Terebratulina australis DW 1520 21°25'S, 175°03'W 447-450 m Terebratulina australis Annuloplatidia curiosa DW 1521 21°19'S, 175°01'W 225-233 m Minutella minuta Basiliola lucida Terebratulina australis Terebratulina japonica DW 1523 21°18'S, 175°00'W 300-302 m Terebratulina reevei Septicollarina zezinae Campages ovalis Terebratulina australis DW 1524 21°17'S, 175°00'W 351-354 m Terebratulina japonica Terebratulina australis CP 1525 21°17'S, 174°59'W 349-351 m Campages ovalis Basiliola lucida Basiliolella colurnus DW 1532 21°44'S, 175°20'W 322 m Terebratulina australis Campages ovalis Basiliola lucida CP 1533 21°44'S, 175°20'W 322-329 m Terebratulina australis Basiliola lucida Basiliolella colurnus DW 1534 21°43'S, 175°19'W 302-327 m Xenobrochus rotundus Terebratulina australis Neoancistrocrania norfolki DW 1535 21°43'S, 175°18'W 268 m Basiliola lucida Campages ovalis Basiliola lucida Basiliolella colurnus DW 1536 21°45'S, 175°21'W 320-323 m Terebratulina australis Campages ovalis

1

606 Bitner M.A.

Station Location Depth Species

Tonga Islands Basiliola lucida Xenobrochus rotundus DWBORDAU 1537 2 21°41'S, 175°19'W 391-421 m Terebratulina australis CampagesBasiliola beecheri ovalis DW 1509 21°05'S, 175°22'W 456-510 m DW 1538 21°39'S, 175°19'W 471-508 m BasiliolaTerebratulina beecheri japonica Basiliola beecheri CP 1539 21°37'S, 175°19'W 558-586 m Basiliola beecheri CP 1510 21°05'S, 175°23'W 461-497 m Terebratulina australis Terebratulina australis Thecidellina maxilla DW 15401514 21°15'S,21°18'S, 175°14'W175°05'W 317130-329-133 m Campages ovalis Basiliola lucida CP 1541 21°15'S, 175°14'W 319-333 m Basiliolella colurnus Terebratulina australis DW 1516 21°21'S, 175°02'W 229-246 m Terebratulina japonica BasiliolaTerebratulina beecheri reevei DW 1543 21°16'S, 175°18'W 427-436 m BasiliolellaSepticollarina colurnus zezinae BasiliolaTerebratulina beecheri australis DW 15441517 21°18'S,21°21'S, 175°18'W175°07'W 441342-443 m m XenobrochusCampages ovalis rotundus Basiliola lucidabeecheri CPDW 1545 1518 21°17'S,21°21'S, 175°17'W175°07'W 444336-447-347 m BasiliolellaTerebratulina colurnus australis CPDW 1546 1520 21°18'S,21°25'S, 175°18'W175°03'W 430447-441-450 m BasiliolaTerebratulina beecheri australis BasiliolaAnnuloplatidia beecheri curiosa DW 1548 20°38'S, 175°03'W 476-478 m DW 1521 21°19'S, 175°01'W 225-233 m TerebratulinaMinutella minuta japonica DW1552 20°38'S, 174°58'W 491-500 m XenobrochusBasiliola lucida rotundus Terebratulina australis Basiliola beecheri Terebratulina japonica DW 1554 20°38'S, 174°58'W 482-498 m Terebratulina japonica DW 1523 21°18'S, 175°00'W 300-302 m Terebratulina reevei Fallax neocaledonensis Septicollarina zezinae BasiliolaCampages beecheri ovalis DW 1555 20°11'S, 174°45'W 591 m Terebratulina japonica Terebratulina australis DW 1524 21°17'S, 175°00'W 351-354 m BasiliolaTerebratulina beecheri japonica CH 1557 20°10'S, 174°42'W 578 m Terebratulina japonica Terebratulina australis CP 1525 21°17'S, 174°59'W 349-351 m CP 1560 19°52'S, 174°39'W 365-372 m AmphithyrisCampages ovalis buckmani CP 1562 19°52'S, 174°42'W 417-424 m AmphithyrisBasiliola lucida buckmani Basiliolella colurnus CPDW 1566 1532 21°02'S,21°44'S, 175°18'W175°20'W 530322-531 m m LeptothyrellaTerebratulina fijiensisaustralis TerebratulinaCampages ovalis reevei DW 1567 21°02'S, 175°19'W 351-356 m ThecidellinaBasiliola lucida maxilla CP 1533 21°44'S, 175°20'W 322-329 m BasiliolaTerebratulina beecheri australis DW 1569 21°02'S, 175°19'W 433 m TerebratulinaBasiliola lucida japonica FrenulinaBasiliolella sanguinolenta colurnus DW 1534 21°43'S, 175°19'W 302-327 m DW 1570 21°02'S, 175°19'W 533-578 m TerebratulinaXenobrochus rotundus japonica Terebratulina australis CP 1578 19°42'S, 174°25'W 329-331 m Amphithyris buckmani Neoancistrocrania norfolki CPDW 1582 1535 18°41'S,21°43'S, 174°03'W175°18'W 79268-82 m m TerebratulinaBasiliola lucida japonica Campages ovalis Basiliola lucida DW 1583 18°37'S, 174°03'W 327-360 m CampagesBasiliola lucida ovalis Basiliolella colurnus DW 15153684 18°36'S,21°45'S, 174°01'W175°21'W 439320 -323m m Terebratulina australisjaponica DW 1585 18°33'S, 173°57'W 578 m TerebratulinaCampages ovalis japonica Basiliola lucida DW 1587 18°37'S, 173°54'W 309-400 m 1Campages ovalis Basiliola lucida DW 1589 18°39'S, 173°54'W 281 m Basiliolella colurnus

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Recent brachiopods from the Tonga Islands, SW Pacific 607

Station Location Depth Species

TongaCP 1593 Islands 19°06'S, 174°18'W 436-442 m Basiliola beecheri

BORDAU 2 Basiliola beecheri DW 1597 20°40'S, 174°55'W 598-610 m Terebratulina japonica Basiliola beecheri DW 1509 21°05'S, 175°22'W 456-510 m DW 1602 20°49'S, 174°57'W 263-320 m TerebratulinaNeoancistrocrania japonica norfolki Basiliola beecherilucida CP 1510 21°05'S, 175°23'W 461-497 m TerebratulinaBasiliolella colurnus australis DW1604 22°16'S, 175°17'W -350 m Terebratulina japonica Thecidellina maxilla DW 1514 21°18'S, 175°05'W 130-133 m Campages ovalis Basiliola lucida Neoancistrocrania norfolki DW 1605 22°17'S, 175°16'W 441 m BasiliolellaAnnuloplatidia colurnus curiosa DW 1516 21°21'S, 175°02'W 229-246 m Terebratulina japonica TerebratulinaBasiliolella colurnus reevei DW 1606 22°16'S, 175°20'W 313-316 m SepticollarinaTerebratulina australiszezinae Campages ovalis Terebratulina australis DW 1517 21°21'S, 175°07'W 342 m DW 1607 22°15'S, 175°23'W 356-367 m CampagesDallina triangularis ovalis DW 1611 23°00'S, 175°47'W 278-323 m BasiliolaTerebratulina lucida australis DW 1518 21°21'S, 175°07'W 336-347 m Terebratulina australis Basiliolella colurnus DW 1520 21°25'S, 175°03'W 447-450 m Terebratulina australis DW 1612 23°02'S, 175°47'W 327-342 m Amphithyris buckmani Annuloplatidia curiosa Campages ovalis DW 1521 21°19'S, 175°01'W 225-233 m Minutella minuta Basiliola lucida Basiliola lucida DW 1614 23°02'S, 175°51' 429-549 m Basiliola beecheri Terebratulina australis Terebratulina australis Terebratulina japonica DW 1523 21°18'S, 175°00'W 300-302 m TerebratulinaBasiliola beecheri reevei DW 1615 23°03'S, 175°53'W 482-504 m SepticollarinaTerebratulina japonicazezinae CampagesAnnuloplatidia ovalis curiosa Terebratulina australisjaponica DW 15241616 21°17'S,23°04'S, 175°00'W175°54'W 351664-354-781 m TerebratulinaAnnuloplatidia japonica curiosa TerebratulinaBasiliola beecheri australis CP 1525 21°17'S, 174°59'W 349-351 m CampagesDyscolia johannisdavisi ovalis DW 1617 23°03'S, 175°53'W 483-531 m Terebratulina japonica Basiliola lucida Fallax neocaledonensis Basiliolella colurnus DW 1532 21°44'S, 175°20'W 322 m TerebratulinaBasiliola beecheri australis DW 1618 24°13'S, 176°18'W 627-656 m CampagesTerebratulina ovalis japonica Basiliola lucidabeecheri CPDW 1533 1619 21°44'S,24°16'S, 175°20'W176°20'W 322591-329-593 m Terebratulina australisjaponica CP 1620 24°18'S, 176°20'W 572 m BasiliolaTerebratulina lucida japonica Basiliolella colurnus DWCH 1 1534621 21°43'S,24°19'S, 175°19'W176°23'W 302570-327-573 m XenobrochusTerebratulina rotundus japonica CP 1625 23°28'S, 176°22'W 824 m TerebratulinaAbyssothyris wyvillei australis NeoancistrocraniaTerebratulina australis norfolki DW 15351628 21°43'S,23°22'S, 175°18'W176°18'W 268400 -416m m BasiliolaAmphithyris lucida buckmani Campages ovalis Basiliola lucida BasiliolellaTerebratulina colurnus australis DW 15361630 21°45'S,23°23'S, 175°21'W176°18'W 320360-323 m m TerebratulinaAbyssothyris wyvillei australis CampagesAmphithyris ovalis buckmani Terebratulina australis DW 1631 23°23'S, 176°18'W 407-443 m 1Amphithyris buckmani Terebratulina australis DW 1632 23°22'S, 176°18'W 613-618 m Terebratulina reevei

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608 Bitner M.A.

Station Location Depth Species

Tonga Islands Basiliola lucida Basiliolella colurnus DW 1634 21°45'S, 175°20'W 321-322 m BORDAU 2 Terebratulina australis CampagesBasiliola beecheri ovalis DW 1509 21°05'S, 175°22'W 456-510 m BasiliolaTerebratulina lucida japonica DW 1635 21°44'S, 175°20'W 320-323 m TerebratulinaBasiliola beecheri australis CP 1510 21°05'S, 175°23'W 461-497 m CampagesTerebratulina ovalis australis DW 1514 21°18'S, 175°05'W 130-133 m BasiliolaThecidellina lucida maxilla DW 1636 21°44'S, 175°20'W 321-331 m Terebratulina australis CampagesBasiliola lucida ovalis Basiliolella colurnus DW 16401516 21°09'S,21°21'S, 175°24'W175°02'W 564229-569-246 m Terebratulina japonica Terebratulina reevei Septicollarina zezinae Terebratulina australis DW 1517 21°21'S, 175°07'W 342 m Campages ovalis Basiliola lucida DW 1518 21°21'S, 175°07'W 336-347 m Terebratulina australis DW 1520 21°25'S, 175°03'W 447-450 m Terebratulina australis Annuloplatidia curiosa DW 1521 21°19'S, 175°01'W 225-233 m Minutella minuta Basiliola lucida Terebratulina australis Terebratulina japonica DW 1523 21°18'S, 175°00'W 300-302 m Terebratulina reevei Septicollarina zezinae Campages ovalis Terebratulina australis DW 1524 21°17'S, 175°00'W 351-354 m Terebratulina japonica Terebratulina australis CP 1525 21°17'S, 174°59'W 349-351 m Campages ovalis Basiliola lucida Basiliolella colurnus DW 1532 21°44'S, 175°20'W 322 m Terebratulina australis Campages ovalis Basiliola lucida CP 1533 21°44'S, 175°20'W 322-329 m Terebratulina australis Basiliola lucida Basiliolella colurnus DW 1534 21°43'S, 175°19'W 302-327 m Xenobrochus rotundus Terebratulina australis Neoancistrocrania norfolki DW 1535 21°43'S, 175°18'W 268 m Basiliola lucida Campages ovalis Basiliola lucida Basiliolella colurnus DW 1536 21°45'S, 175°21'W 320-323 m Terebratulina australis Campages ovalis

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